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Author contributions: Kim JH, Ha HL, Son HY and Yu DY planned the experiments and analyses; Kim JH, Ha HL, Shin HJ, Park YH, Kwon TH, Jung MR, Moon HB, Cho ES, Son HY and Yu DY performed the experiments and analyzed the results; Kim JH and Yu DY wrote the manuscript; all the authors discussed the results and commented on the manuscript.

Supported by The World Class Institute (WCI) Program of the National Research Foundation of Korea (NRF), No. WCM0101222; Ministry of Education, Science and Technology of Korea (MEST), the National R and D Program for Cancer Control, Ministry of Health and Welfare, South Korea, No. BCM0061213; KRIBB Research Initiative Program Grant, No. KGM3141312